View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Digital Repository @ Iowa State University Ecology, Evolution and Organismal Biology Publications Ecology, Evolution and Organismal Biology 2020 Exotic species drive patterns of plant species diversity in 93 restored tallgrass prairies Andrew D. Kaul Iowa State University Brian J. Wilsey Iowa State University, [email protected] Follow this and additional works at: https://lib.dr.iastate.edu/eeob_ag_pubs Part of the Biodiversity Commons, Ecology and Evolutionary Biology Commons, Natural Resources and Conservation Commons, Plant Sciences Commons, and the Soil Science Commons The complete bibliographic information for this item can be found at https://lib.dr.iastate.edu/ eeob_ag_pubs/440. For information on how to cite this item, please visit http://lib.dr.iastate.edu/howtocite.html. This Article is brought to you for free and open access by the Ecology, Evolution and Organismal Biology at Iowa State University Digital Repository. It has been accepted for inclusion in Ecology, Evolution and Organismal Biology Publications by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Exotic species drive patterns of plant species diversity in 93 restored tallgrass prairies Abstract A primary goal of restoration ecology is to understand the factors that generate variability in species diversity and composition among restorations. Plant communities may assemble deterministically toward a common community type, or they may assemble stochastically, ending differently because of weather conditions during establishment, soil legacy effects, or exotic species propagule pressure. To test these alternative hypotheses, we sampled plant communities and soil at 93 randomly selected restored prairies distributed throughout Iowa, USA. Five remnant sites were sampled as a reference. We tested our hypotheses using multiple regressions and investigated the strength of direct and indirect effects on species diversity and richness using structural equation models. The prairie restorations were highly variable in their age, size, diversity, soil characteristics, and how they were managed post‐seeding. The strongest predictor of plant species richness and diversity was the degree of invasion, as measured by the abundance of exotic species. Restorations planted with species‐rich seed mixes had reduced exotic species abundance, which led indirectly to higher species richness of restorations. Sites with higher organic matter and a more linear shape had a direct positive effect on exotic abundance, which in turn decreased diversity. We found little support for deterministic assembly, and diversity did not increase with the age of planting. Our results indicate that restored prairie communities tend to assemble into states of high or low diversity, driven by invasion from exotic plant species. Management of exotic species is essential for maximizing species diversity in temperate grassland restorations. Keywords grassland, invasive species, prairie, restoration, plants, tallgrass Disciplines Biodiversity | Ecology and Evolutionary Biology | Natural Resources and Conservation | Plant Sciences | Soil Science Comments This article is published as Kaul, Andrew D., and Brian J. Wilsey. "Exotic species drive patterns of plant species diversity in 93 restored tallgrass prairies." Ecological Applications (2020): e2252. doi: 10.1002/ eap.2252. Copyright by the Ecological Society of America. Posted with permission. This article is available at Iowa State University Digital Repository: https://lib.dr.iastate.edu/eeob_ag_pubs/440 Ecological Applications, 0(0), 2020, e02252 © 2020 by the Ecological Society of America Exotic species drive patterns of plant species diversity in 93 restored tallgrass prairies 1 ANDREW D. KAUL , AND BRIAN J. W ILSEY Department of Ecology Evolution and Organismal Biology, Iowa State University, 251 Bessey Hall, 2200 Osborn Drive, Ames, Iowa 50011 USA Citation: Kaul, A. D., and B. J. Wilsey. 2020. Exotic species drive patterns of plant species diversity in 93 restored tallgrass prairies. Ecological Applications 00(00):e02252. 10.1002/eap.2252 Abstract. A primary goal of restoration ecology is to understand the factors that generate variability in species diversity and composition among restorations. Plant communities may assemble deterministically toward a common community type, or they may assemble stochasti- cally, ending differently because of weather conditions during establishment, soil legacy effects, or exotic species propagule pressure. To test these alternative hypotheses, we sampled plant communities and soil at 93 randomly selected restored prairies distributed throughout Iowa, USA. Five remnant sites were sampled as a reference. We tested our hypotheses using multiple regressions and investigated the strength of direct and indirect effects on species diversity and richness using structural equation models. The prairie restorations were highly variable in their age, size, diversity, soil characteristics, and how they were managed post-seeding. The strongest predictor of plant species richness and diversity was the degree of invasion, as measured by the abundance of exotic species. Restorations planted with species-rich seed mixes had reduced exotic species abundance, which led indirectly to higher species richness of restorations. Sites with higher organic matter and a more linear shape had a direct positive effect on exotic abun- dance, which in turn decreased diversity. We found little support for deterministic assembly, and diversity did not increase with the age of planting. Our results indicate that restored prairie communities tend to assemble into states of high or low diversity, driven by invasion from exo- tic plant species. Management of exotic species is essential for maximizing species diversity in temperate grassland restorations. Key words: grassland; invasive species; prairie; restoration; plants; tallgrass. reproducing this high diversity in restorations remain INTRODUCTION understudied (Symstad and Jonas 2011). A fundamental question in restoration ecology is Two theories make distinct predictions on how species whether the full biodiversity represented in reference assemble into diverse communities over time during areas can be achieved in restored areas, given enough restoration (reviewed by Temperton and Hobbs 2004). time (Bradshaw 1987). Biodiversity is a key integrative The deterministic model predicts that communities will measure that has been linked to higher ecosystem func- assemble toward a predictable and consistent commu- tions in grasslands, including productivity, nitrogen min- nity composition and diversity over time as determined eralization, and litter decomposition (Hooper et al. by local environmental conditions. Many restoration 2005, Isbell et al. 2011). It is also positively associated seed mixes include later successional species at establish- with stability, especially the consistency of biomass pro- ment to overcome dispersal limitation and bypass the duction across years (Tilman et al. 2001, 2006, Isbell early colonization stage by design. Under the determinis- et al. 2009), and resistance to extreme precipitation tic model, the composition of the seed mix, time, and events (Isbell et al. 2015). Biodiversity is also important local environmental conditions would be the most to pollinators in grasslands, because animal-pollinated important variables for predicting restoration outcomes, forbs are more abundant in more diverse sites (Haddad with diversity generally expected to increase over time as et al. 2001). Remnant prairies can have 15–20 species in species recruit from the mix (Fig. 1; Wilsey and Martin areas <0.5 m2 (Partel¨ and Zobel 1999, Martin et al. 2015). However, few studies have tested this assumption, 2005, Polley et al. 2005), and restoring this high diversity and some find that diversity peaks after a few years and remains a challenge. The factors that are responsible for then declines over time as tall grasses dominate (Pimm 1991, Sluis 2002, Grman et al. 2013). / Manuscript received 11 December 2019; revised 9 July 2020; The stochastic alternate state model predicts that fac- accepted 16 August 2020. Corresponding Editor: Amy J. Sym- tors that are somewhat random set the trajectory of stad. community development, which can lead to alternate 1E-mail: [email protected] Article e02252; page 1 Ecological Applications Article e02252; page 2 ANDREW D. KAUL AND BRIAN J. WILSEY Vol. 0, No. 0 FIG. 1. Conceptual relationships between community assembly models and predictors of restoration outcomes in prairie grass- lands. Soil organic matter is abbreviated as SOM. compositions. For example, weather during initial years affected by species sorting along abiotic gradients, dis- post-seeding can lead to unpredictable outcomes if con- persal, variation in assembly history (Questad and Fos- ditions are far from the climatic norm (Manning and ter 2008, Martin and Wilsey 2012), and actions by Baer 2018, Groves and Brudvig 2019), and compositions animals such as urine deposition or grazing (Steinauer that form during this time can have long-term impacts and Collins 1995, Loreau 2000, Chase 2003, Soininen through priority effects. Soil legacy or priority effects et al. 2007, Martin and Wilsey 2015). Restoring beta can have strong influences on assembly, resulting in dif- diversity is extremely important for grassland